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Aspects of the Ecology of Bartramia Halleriana and Crossocalyx Hellerianus in Oceanic Deciduous Woodland

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Aspects of the Ecology of Bartramia Halleriana and Crossocalyx Hellerianus in Oceanic Deciduous Woodland bioRxiv preprint doi: https://doi.org/10.1101/2020.04.28.066522; this version posted April 30, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Aspects of the ecology of Bartramia halleriana and Crossocalyx hellerianus in oceanic deciduous woodland Des A. Callaghana and Jamie Bevanb aBryophyte Surveys Ltd, Bristol, UK; bNatural Resources Wales, Neath, UK Correspondence to: Des A. Callaghan, 20 Thornleigh Road, Bristol, BS7 8PH, UK. E-mail: [email protected] Abstract Introduction: This study investigates the ecology of two boreal-montane bryophytes, the saxicolous moss Bartramia halleriana and the epixylic liverwort Crossocalyx hellerianus, at the edge of their ranges, in oceanic deciduous woodland. Methods: The study site comprises two adjoining woodlands, Allt Penyrhiw-iar (canopy trees ca. 100 yr old) and Allt Rhyd Groes (ca. 200 yr), Carmarthenshire, UK. The distribution and abundance of B. halleriana and C. hellerianus was surveyed, relevés sampled to record habitat and community composition, and sporophyte frequency and stage of development measured. Light climate of B. halleriana was investigated via hemispherical photography, and abundance of large rotten logs used as a measure of habitat quality for C. hellerianus. Results and discussion: Four subpopulations of B. halleriana occur, comprising 21 individual- equivalents (occupied 1 m grid cells), all on mildly base-rich mudstone of north-facing rockfaces, with very little direct solar radiation and a diverse assemblage of bryophytes. Sporophyte are scarce. A total of 143 individual-equivalents (occupied logs or trees) of C. hellerianus was recorded, an exceptional population in Wales. Most of the population is within Allt Rhyd Groes, where large rotten logs are much more abundant due to greater woodland age. The liverwort mainly occupied the sides of decorticated rotten logs, amongst a sparse community composed mainly of other small liverworts and Cladonia lichens. Neither sporophytes nor perianths were found. Further research could usefully focus on the description and measurement of favourable habitat for each species and associated physiological tolerances. Keywords: moss, liverwort, abundance, light climate, sporophyte frequency, deadwood. 1 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.28.066522; this version posted April 30, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Introduction Sites that are rich in rare and scarce species are of special conservation value and there is a clear need to ensure their protection and effective management. Indeed, criteria used to select sites for safe-guarding commonly include reference to such species, for example regarding ‘restricted- range species’ in the context of ‘key biodiversity areas’ (Eken et al. 2004). In Britain, the statutory nature conservation agencies have a duty under the Wildlife and Countryside Act 1981 (as amended), to notify any area of land which in their opinion is 'of special interest by reason of any of its flora, fauna, or geological or physiographical features' (Bainbridge et al. 2013). These areas are known as Sites of Special Scientific Interest (SSSIs). Some of the criteria used for the identification of SSSIs are based on a scoring system, with thresholds to determine occurrence within sites of nationally important assemblages of rare and scarce species, including bryophytes (Bosanquet et al. 2018). Due to their national scarcity (Pescott 2016), Bartramia halleriana Hedw. (Bartramiales: Bartramiaceae) and Crossocalyx hellerianus (Nees ex Lindenb.) Meyl (Jungermanniales: Anastrophyllaceae) can contribute to scores used for the identification of SSSIs in Britain, and as a consequence they can become a focus of site-based monitoring and management activities, for which knowledge of their ecology is fundamental. The monoicous moss B. halleriana is a boreal-montane species (Hill and Preston 1998), occurring frequently in some parts of Europe and Asia, and very rarely in North America, Central America and Africa (Blockeel et al. 2014; Fransén 2004). In Britain, it was recorded in 57 10 km grid cells during 1990–2013, in northern England, Scotland and Wales (Blockeel et al. 2014). The dioicous liverwort C. hellerianus also exhibits a boreal-montane distribution pattern (Hill and Preston 1998), essentially circumboreal and largely continental, occurring in Europe, Asia and North America (Blockeel et al. 2014; Hong 1996; Sabovljevic et al. 2019; Schuster 1969; Schill and Long 2003). Its range in Britain is similar to B. halleriana, being recorded in 38 10 km grid cells during 1990–2013, confined to northern England, Scotland and Wales (Blockeel et al. 2014). To date, no research has been undertaken regarding the ecology of either species in Britain. The purpose of this study is to investigate such, to help inform conservation decisions and research priorities. Method Study area The study site (52°07'05"N, 3°48'37"W; 160–350 m a.s.l.) comprises two contiguous woodland compartments, Allt Penyrhiw-iar (17.6 ha) and Allt Rhyd Groes (14.6 ha), Carmarthenshire (v.-c. 2 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.28.066522; this version posted April 30, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 44), South Wales. It supports a nationally important assemblage of rare and scarce bryophytes of woodland, including B. halleriana and C. hellerianus. The site is within Allt Rhyd y Groes National Nature Reserve (NNR), and the much larger protected areas of Cwm Doethie–Mynydd Mallaen SSSI and Cwm Doethie–Mynydd Mallaen Special Area of Conservation (SAC). The woodland mainly comprises W17 Quercus petraea–Betula pubescens–Dicranum majus woodland and W11 Quercus petraea–Betula pubescens–Oxalis acetosella woodland (Rodwell 1991), located on steep hillsides, with geology dominated by mudstones, weathering to produce clay soils. The climate is oceanic, with 176 rain-days yr-1 (days with >1 mm rain) and average temperatures of 14.6°C during the hottest month (July) and 3.3°C during the coldest month (February) for the period 1961–2002 (Met Office data supplied through the UK Climate Impact Programme). The NNR was established in 1959 and has since been managed for nature conservation, with a general policy of low intervention woodland management. Conservation management has recently included thinning of understorey trees and shrubs in some parts and controlled light grazing by sheep (ca. 0.05 LSU ha-1 yr-1) is practised throughout. Little is known of the long-term management history of the two woodland compartments within the study area, though in 1821, 249 oaks from ‘Allt Rhyd y Groes’ were sold by Cawdor Estate, the landowner at that time, possibly for ship building (NRW 2020). Indeed, canopy oak trees in both compartments are relatively even-aged, indicating clear-felling in the past, perhaps followed by re-planting. However, those in Allt Rhyd Groes average ca. 200 yr old, while those in Allt Penrhiw-iar are mostly <100 yr (NRW 2020). Both woodlands have been identified as ‘ancient woodland’ (Lister and Whitebread 1988) and are likely to have a long history of tree cover, notwithstanding episodes of felling for timber. Taxonomy Taxonomy follows Hodgetts et al. (2020) for bryophytes, Smith et al. (2009) for lichens and Stace (2010) for vascular plants. Geographic reference system Geographic coordinates follow the Ordnance Survey (OS) National Grid reference system (EPSG: 27700). Grid cells are referred to by the coordinates of the south-west corner. Distribution and abundance COSWIC (2011) defined an individual of B. halleriana as “a discrete colony (clump or tuft of moss consisting of many shoots)”. Following trial in the field, including further definition of “discrete” 3 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.28.066522; this version posted April 30, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. (≥10 cm distance from any other count unit) and “many shoots” (≥10), this was not followed presently because it ignored frequent occurrences of the moss that did not fit within the definition. Indeed, whilst B. halleriana is a moss with a ‘tuft’ growth form (Hill et al. 2007), colonies frequently have fuzzy boundaries, for example with tufts sometimes blending into one another, larger tufts sometimes becoming fragmented due to disturbance events, and other occurrences not being tufts at all, but rather scattered shoots amongst other mosses, making the counting of discrete colonies troublesome. Instead, an ‘individual-equivalent’ of B. halleriana was here defined as an occupied 1 m grid cell, following Bergamini et al. (2019). Such were counted after marking- out locations of the moss in the field, and where necessary using a tape-measure to determine number of occupied 1 m grid cells. An individual-equivalent of C. hellerianus is defined as an occupied log or tree, again following Bergamini et al. (2019). An inventory of previously known locations of both species within the study area was compiled from records within the national recording database of the British Bryological Society (BBS), held by the Biological Records Centre (Wallingford, UK), and reports of previous bryophyte surveys (Newton 1997, 2002, 2008). During September 2019 and February–March 2020, the study site was searched for locations of the two species, revisiting all former locations and ensuring good coverage of other areas.
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